Darwiniana ISSN: 0011-6793 [email protected] Instituto de Botánica Darwinion

Wheeler, Gerald A.; Guaglianone, Encarnación R. Notes on south American (): C.camptoglochin and C. microglochin Darwiniana, vol. 41, núm. 1-4, 2003, pp. 193-206 Instituto de Botánica Darwinion Buenos Aires, Argentina

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41(1-4): 193-206. 2003

NOTES ON SOUTH AMERICAN CAREX (CYPERACEAE): C. CAMPTOGLOCHIN AND C. MICROGLOCHIN

GERALD A. WHEELER1 AND ENCARNACIÓN R. GUAGLIANONE 2,3 1University of Minnesota Herbarium. J. F. Bell Museum of Natural History, St. Paul, Minnesota 55108-1095, United States of America. E-mail: [email protected] 2Instituto de Botánica Darwinion. Casilla de Correo 22, B1642HYD, San Isidro, Buenos Aires, Argentina. E-mail: [email protected]

ABSTRACT: Wheeler, G. A. & Guaglianone, E. R. 2003. Notes on South American Carex (Cyperaceae): C. camptoglochin and C. microglochin. Darwiniana 41(1-4): 193-206.

Carex microglochin (Cyperaceae), a member of Carex sect. Leucoglochin, is a bipolar species that, for the most part, occurs in the northern part of the Northern Hemisphere and in the Andean region of . A majority of past and present workers have treated the northern as subsp. microglochin, whereas the southern plants have been referred to subsp. fuegina. However, it is demonstrated in this paper that two closely-related yet taxonomically-distinct species, C. camptoglochin and C. microglochin, occur in South America, these differing both morphologically and ecologically. Regarding the South American distribution of these two species, it is documented below that: (1) C. microglochin occurs sporadically throughout nearly the full extent of the Andes; (2) C. microglochin and C. camptoglochin grow sympatrically in both the boreal and austral parts of South America; and (3) C. camptoglochin occurs as a disjunct on the (Islas Malvinas).

Key words: , Cyperaceae, Carex, sect. Leucoglochin, Carex camptoglochin , .

RESUMEN: Wheeler, G. A. & Guaglianone, E. R. 2003. Notas sobre Carex (Cyperaceae) de Sud América: C. camptoglochin y C. microglochin. Darwiniana 41(1-4): 193-206.

Carex microglochin (Cyperaceae, Carex, sect. Leucoglochin) es una especie bipolar. La mayoría de los autores, han considerado las plantas del hemisferio Norte como subsp. microglochin, mientras que las plantas del hemisferio Sur se refirieron a la subsp. fuegina. En esta nota se demuestra que este último taxón corresponde a dos especies estrechamente relacionadas aunque taxonómicamente distintas, C. camptoglochin y C. microglochin, ambas en Sud América, las que difieren tanto en su morfología como en su hábitat. En relación con la distribución sudamericana de estas dos especies, se documenta que: (1) C. microglochin habita ocasionalmente casi a todo lo largo de los Andes; (2) C. microglochin y C. camptoglochin son simpátricas en Sud América, tanto austral como septentrional, y (3) C. camptoglochin habita como disyunta en las Islas Malvinas (Falkland Islands).

Palabras clave: Taxonomía, Cyperaceae, Carex, sect. Leucoglochin, Carex camptoglochin, Carex microglochin. INTRODUCTION It is well known, and of phytogeographical microglochin Wahlenb., particularly its relation- interest, that some plants occur in both North and ship to C. camptoglochin V. I. Krecz., whose type South America though with varying degrees of comes from austral South America. A review of disjunction (Raven, 1963). Contrary views exist for apposite literature indicates that a majority of past some amphitropical taxa, however, such as the and present authors (e.g., Kükenthal, 1909; Barros, taxonomic status and bipolar distribution of Carex 1947, 1969; Moore, 1968, 1974, 1983; Moore & 3 Miembro de la Carrera del Investigador, CONICET

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Chater, 1971) maintain that C. microglochin subsp. and Southern Hemispheres. Moore & Chater (1971) microglochin occurs only in the Northern treated all such plants from the Northern. Hemisphere, whereas all South American material is Hemisphere as C. microglochin (their subsp. assignable to subsp. fuegina Kük. But contrarily, microglochin), whereas all such plants from South the present study supports the claim of other America were referred to C. camptoglochin (their C. workers (Roivainen, 1954; Kukkonen, 1970) who microglochin subsp. fuegina). Contrarily, suggest that two closely-related yet taxonomically- Roivainen (1954) suggested that both C. distinct species, C. camptoglochin and C. microglochin and C. camptoglochin (his C. microglochin, occur in South America. Indeed, it is oligantha) occur in , with the shown below that these two species, which differ former growing in the drier, eastern parts of Fuegia both morphologically and ecologically, grow and the latter in the wetter, western and sympatrically in both the far northern and southern southwestern parts. A subsequent study by parts of South America. Moreover, we document Kukkonen (1970) supports Roivainen’s claim that that C. microglochin occurs sporadically C. camptoglochin and C. microglochin are distinct throughout nearly the full length of the Andes and, species, as does also the mention of both names by further, that C. camptoglochin grows as a disjunct Reznicek (1990). Nevertheless, a great majority of on the Falkland Islands (Islas Malvinas). South American authors (e.g., Barros, 1947, 1969; Both C. camptoglochin and C. microglochin, as Marticorena & Quezada, 1985; Guaglianone, 1996) well as the austral South American endemic C. have followed Kükenthal (1909) and Moore & kingii (Boott) Reznicek, belong in Carex sect. Chater (1971), the latter who claim that a large Leucoglochin Dumort. A key to the three South overlap in characters precludes recognition of both American members of this section is given near the taxa in Tierra del Fuego. It is also worth noting here end of this report. For essentially the remainder of that most authors, while discussing the taxonomic this paper, however, C. kingii is not discussed status of the microglochin-like plants that occur in further and the term “microglochin-like” is restric- Tierra del Fuego, offer no opinion as to the status of ted to C. camptoglochin and C. microglochin, the similar populations that grow elsewhere on the only two of the three that possess strongly deflexed South American continent. perigynia at maturity and a rachilla that lacks a hook-like appendage at the apex. MORPHOLOGY Moore & Chater (1971: 328) studied nine BRIEF HISTORICAL OUTLINE characters of the microglochin-like plants that Carex microglochin, whose type comes from occur in South America and concluded that none of northern Lapland in Scandinavia, is well known those characters could be used to positively from North America and parts of Europe and Asia, differentiate the two taxa in question. The cha- though on the North American continent it is racters they studied were: stem length, stem reported only as far south as south-central Colora- diameter, length of longest leaf, number of male do (Mackenzie, 1931–1935; Hermann, 1970). It is flowers, number of female flowers, length of largest unknown from Central America (Chater, 1994) but perigynium, width of largest perigynium, emergent has been reported from (Mora & Rangel, length of largest rachillae, and stipe length. But as is 1983; Luteyn, 1999), (Jørgensen & Ulloa, evident in Table 1 and in the key given near the end 1994; Luteyn, 1999), and Peru (Luteyn, 1999) in of this report, there are several other characters, not northern South America. The type of C. utilized by these workers, that provide clear-cut camptoglochin, on the other hand, comes from Ba- differences between the two taxa. Salient morpho- hía Orange on Isla Hoste in southern Tierra del logical differences are discussed immediately Fuego. This was originally called C. oligantha below. Boott (1867), but Kreczetowicz (1937) created a new The pistillate scales of the two taxa differ name for it, as the former name was a later homonym. considerably in both size and shape. Those of C. In general, two different distribution scenarios camptoglochin (Fig. 1) are lanceolate and longer have been presented in the literature regarding the than the perigynium, with the apex acute to acumi- microglochin-like plants that occur in the Northern nate and the broad midrib frequently projecting

194 G. A. WHEELER & E. R. GUAGLIANONE. Notes on South American Carex: C. camptoglochin and C. microglochin

Table 1.- A selected morphological comparision of Carex camptoglochin and C. microglochin in South America.

Character C. camptoglochin C. microglochin

Growth habit decumbent old stems creeping sympodial rhizomes Rhizome/decumbent stem width (mm) 0.9 - 1.4 0.4 - 0.7 Number of male flowers per spike 2 - 3 5 - 6 Anther length (mm) 1.6 - 2.4 0.8 - 1.6 Number of female flowers per spike 2 - 5 (most often 3 or 4) 3 - 12 Lowest pistillate scale length (mm) 6 - 12 (-15) 2 - 4 Pistillate scale shape lanceolate ovate Pistillate scale apex acute to acuminate or awned obtuse to subacute Pistillate scale color reddish brown pale brown to brown Perigynium length (mm) 4.6 - 8.2 2.8 - 4.8 Perigynium stipe type and length (mm) long-stipitate, 0.7 - 2 short-stipitate, < 0.7 Perigynium stipe shape when deflexed at maturity inverted U-shaped flattened-thickish Perigynium beak color reddish brown stramineous or brownish Number of vascular bundles in rachilla 2 1 Achene length (mm) 2.5 - 3.4 2 - 2.5 beyond the apex as a smooth awn; in contrast, C. that, in general, the perigynia of C. camptoglochin microglochin (Fig. 2) has ovate scales that are are longer than those of C. microglochin, though shorter than the perigynium, with the apex obtuse to this character by itself, as pointed out by Moore & subacute and the comparatively narrow midrib Chater (1971), cannot be used to positively separate seldom reaching the tip. Moreover, the lowermost the two species as some overlap in length does pistillate scale of the former species sometimes occur. extends to, or even exceeds, the tip of the exserted As indicated in Table 1 and in the key further rachilla, which is never the case in the latter. In below, Carex camptoglochin has, generally regard to the staminate scales, it is worthy of note speaking, fewer perigynia per spike than C. that C. camptoglochin has 2 or 3 per spike, while C. microglochin. Also, both the achenes and anthers microglochin has 5 or 6. Also, both the staminate of C. camptoglochin are slightly longer than the and pistillate scales of C. camptoglochin are same features in C. microglochin. All three of these reddish brown, whereas those of C. microglochin characters are useful in identification, with the last are pale brown to brown. being most useful at anthesis and the first two when Some characteristics of the perigynia also perigynia are mature. differentiate the two species. For instance, the base As first pointed out by Kukkonen (1970: 144), of the perigynium in C. camptoglochin is long- “even the mode of growth is different in these two stipitate, whereas C. microglochin has a short species” (see Figs. 1, 2). It is usually reported stipe. In the former, when plants are mature and correctly (e.g., Kükenthal, 1909; Mackenzie, 1931– perigynia usually deflexed, the stipe is strongly 1935) that C. microglochin is rhizomatous, with the inverted U-shaped (Fig. 1); contrarily, in the latter plants producing creeping sympodial rhizomes. On species, though the perigynia are also usually the other hand, C. camptoglochin is unusual, deflexed at maturity, the stipe is flattened-thickish though not unique in Carex, in having the next with the perigynia appearing subsessile on the year’s shoots, both vegetative and flowering, arise rachis (Fig. 2). Moreover, the perigynium beak of C. from dormant buds in the axils of dried leaves of camptoglochin is reddish brown near the apex, prolonged, decumbent old stems, with the terminal whereas in C. microglochin the apex of the beak is and upper nodes producing mostly newly- stramineous or brownish. It is also worth noting developed flowering shoots and the lower nodes

195 Darwiniana 41(1-4). 2003

Fig. 1.- Carex camptoglochin. A: plant. B: flattened apex of leaf. C: leaf with sheath and ligule. D: spike. E: scale. F: mature spike with perigynia deflexed, showing the long stipes strongly inverted U-shaped. G: transection of perigynium with rachilla and trigonous achene. H: achene, with part of the style. A, B: from T.B.P.A. 1732; C, F, G, H: from Goodall 2417; D, E: from Biganzoli 744. vegetative shoots. Moreover, the roots of C. Lastly, but notably, Kukkonen (1970) reported camptoglochin are produced along the decumbent anatomical differences between these two species. old stems, generally at the lower nodes, whereas After studying a number of characters, both those of C. microglochin are produced at the base quantitative (e.g., presence or absence of prickles or of flowering and vegetative shoots, with essentially papillae, size of stomata, position of vascular none produced along the rhizome itself. Also, the bundles) and qualitative (e.g., position of stomata, slender, elongated creeping rhizomes of C. number of air-cavities, differentiation of microglochin are appreciably narrower than the chlorenchyma), he maintained that both C. chord-like, decumbent stems of C. camptoglochin. camptoglochin and C. microglochin occur in Tierra Regarding growth habit, C. camptoglochin del Fuego. Also, microscopy (by E.R.G.) reveals resembles that of some carices from the Northern that the rachilla of C. camptoglochin has two Hemisphere, such as C. chordorrhiza L. f. and C. vascular bundles, whereas C. microglochin is limosa L. (e.g., Mackenzie, 1931–1935; Damman, reported (Reznicek, 1990) to have only a single bun- 1964; Reznicek & Catling, 1986). dle.

196 G. A. WHEELER & E. R. GUAGLIANONE. Notes on South American Carex: C. camptoglochin and C. microglochin

Fig. 2.- Carex microglochin. A: plant. B, C: spikes in different stages of anthesis. D: mature spike with perigynia deflexed, lacking long stipes. E: transection of perigynium with rachilla and subtrigonous achene. F: achene and style. A, E, F: from Moore 1343. B: from Notø 31. C: from Kiesling 7996. D: from Cabrera 31756.

GEOGRAPHIC DISTRIBUTION AND summit. Farther south, in Tierra del Fuego, C. ECOLOGICAL REQUIREMENTS microglochin occurs in the north and northeastern Some of the South American microglochin-like portions of Fuegia, while C. camptoglochin grows populations that occur north of 30°S lat. are clearly in the west and southwestern portions. The two assignable to C. microglochin (Fig. 4), as they are species grow sympatrically in southern Fuegia, essentially identical to populations of this species where the elevation is higher, but C. microglochin occurring in the Northern Hemisphere. Within this is unknown from Antártica Chilena Province, , same extensive area C. camptoglochin (Fig. 3) is an area that, generally speaking, lies southwest of known, thus far, only from Colombia and Ecuador. the Andean summit; contrarily, C. camptoglochin Few microglochin-like plants have been seen is well known from this Chilean province (e.g., Isla between 30°S lat. to 45°S lat., but in the southern Hoste, Isla Navarino, Isla Wollaston) as well as from half of C. microglochin is known from Isla de los Estados, an Argentine island which lies both sides of the Andes, whereas C. just off the southeastern tip (Península Mitre) of camptoglochin is confined west of the Andean South America.

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Fig. 3.- Map of South America showing the distribution of Carex camptoglochin; starred circle represents the holotype locality.

198 G. A. WHEELER & E. R. GUAGLIANONE. Notes on South American Carex: C. camptoglochin and C. microglochin

Of phytogeographic interest, the microglochin- Unfortunately, very little information exists in the like plants that occur on the Falkland Islands (Islas literature concerning microenvironmental Malvinas) are assignable to C. camptoglochin, differences between these two species, but perso- these known from “peat bogs” on both large nal correspondence and data on herbarium sheet islands. Notably, there are also several other plants, labels have been most valuable and instructive. such as Agrostis magellanica Lam., Astelia pumila Habitat information for C. camptoglochin and C. (Forster f.) Gaudich., and Cortaderia pilosa microglochin are presented and discussed under (D’Urv.) Hackel that display a similar distribution each species further below. pattern, i.e., the species grow primarily in Sphagnum bogs in southern Fuegia and farther CONCLUSIONS westward, are uncommon or absent from north and If the view put forth by Moore & Chater (1971: northeastern Fuegia, but also are disjunct in peat 330) is correct that “... all Northern Hemisphere ma- bogs on the Falkland Islands (Islas Malvinas) terial be included in ssp. microglochin, while South (Moore, 1983). American plants comprise ssp. fuegina”, then In regard to the climatic requirements of these presumably all microglochin-like plants from South two species in austral South America, there also America should possess features essentially appear to be noteworthy differences. The frequent identical to those of the type of C. camptoglochin occurrence of the bog-loving C. camptoglochin in (their C. microglochin ssp. fuegina). But the western Fuegia and far western Patagonia is not morphological evidence presented above strongly unexpected because of wetter conditions on the contradicts this view, indicating instead that two western side of the Andes due to orographic distinct species, C. camptoglochin and C. precipitation, a result of moisture picked up over the microglochin, occur in South America. Moreover, it Pacific Ocean. The heavy rainfall in the western is documented here that both species occur in portions of Patagonia and Fuegia, concomitant with northern South America, with the specimens of C. cool temperatures, is conducive to the camptoglochin from Colombia and Ecuador (see accumulation of peat deposits, which supports bog specimen citations further below) representing the vegetation. Contrarily, C. microglochin occurs in first authenticated report of this species north of the relatively drier northern and eastern portions of Patagonia. Fuegia, which in large part is due to the rain-shadow Although we consider C. camptoglochin and C. effect of the Andes. As pointed out by Moore microglochin to be distinct species, it would be (1983: 4), “…there is a dramatic rainfall gradient from remiss if, at this point, we failed to mention some south and west Fuegia to the conspicuously more probable reasons why these two taxa have been arid rain shadow of north and east Isla Grande [in confused in the past. Firstly, because the pistillate northeastern Tierra del Fuego]”. In these portions scales of both species are pronouncedly caducous, of Fuegia, C. microglochin grows primarily in moist most (if not all) of the scales are absent when fruits to wet minerotrophic sites. Farther south in Fuegia, are fully mature (i.e., when the perigynia are however, where rainfall is more abundant, C. deflexed) which is, unfortunately, when most camptoglochin and C. microglochin occur collections of these plants are made. Secondly, the sympatrically. strongly-deflexed perigynia are very easily One can presumably speculate that the detached from the inflorescence, consequently, the Sphagnum-dominated bogs, preferred by C. plants typically have few perigynia remaining on camptoglochin, have lower pH values (i.e., they are the rachis when collected. Lastly, habit differences more acidic) than the minerotrophic wet meadows, between these two species are difficult to assess on moist grasslands, and swamps that are often many herbarium sheets because only above- recorded as habitats for C. microglochin. Thus, ground (or above-stratum) fruiting culms have been although C. camptoglochin and C. microglochin collected. In regard to the last point, and certainly are known to grow sympatrically in northern South worth emphasizing, we and other workers (e.g., America as well as in southern Fuegia, it seems Damman, 1964; Jermy et al., 1982; Reznicek & reasonable to suggest that these two species Catling, 1986) strongly urge that when collecting require different edaphic conditions for growth.

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Fig. 4.- Map showing the distribution of Carex microglochin in South America. For the generalized range of this species in the Northern Hemisphere see Moore & Chater (1971: 321, Fig. 1 F).

200 G. A. WHEELER & E. R. GUAGLIANONE. Notes on South American Carex: C. camptoglochin and C. microglochin

Carex (as well as other cyperaceous plants) all brown. Inflorescence a single, terminal andro- flowering and vegetative parts of the plant should gynous spike 7-14 mm long, 6-9(-13) mm wide; be collected, not just fruiting culms. bracts absent. Staminate part 2-3-flowered; scales 3- Notably, every microglochin-like plant examined 5.5 mm long, 0.8-1.6 mm wide, ovate-lanceolate, apex from South America could readily be assigned to acute or mucronate, pale reddish brown, with either C. camptoglochin or C. microglochin, as hyaline margins and greenish center, 1(-3)-veined. propounded in the discussion of morphology Pistillate part 2-5-flowered (most often 3 or 4); scales above and pointed out in Table 1, where the caducous, equaling to much exceeding the differentiating characters between the two species perigynia, 5-12(-15) mm long, 1.6-2.6 mm wide, are summarized. Because it is abundantly clear from lanceolate, apex acute to acuminate or awned, this study that these two taxa differ both morpholo- reddish brown, with narrow hyaline margins and gically and ecologically, it is strongly recommended greenish center, 1-3-veined, with the midrib fre- here that henceforth C. camptoglochin and C. quently excurrent as a smooth awn. Perigynia 4.6- microglochin be considered as distinct species in 8.2 mm long, 0.7-1.1 mm wide, ascending at first but the Carex flora of South America. soon becoming wide spreading or more often deflexed, subulate or linear-lanceolate, suborbicular CAREX CAMPTOGLOCHIN AND C. in cross-section, glabrous, membranaceous, MICROGLOCHIN greenish or pale reddish brown to reddish brown, Carex camptoglochin V. I. Krecz., Bot. Mater. Gerb. weakly to moderately many-striate, the margins Bot. Inst. Komarova Akad. Nauk S. S. S. R., 7: smooth, long-stipitate, with the 0.7-2 mm stipe 34. 1937. Carex oligantha Boott, Ill. Carex 4: strongly inverted U-shaped at maturity, very 174. 1867, non Steudel (1855) et nec Phil. (1891). gradually tapering to the apex; beak not well C. microglochin Wahlenb. subsp. fuegina differentiated from the body, ca. 1.5-2.5 mm long, Kük., Bot. Jahrb. Syst. 27: 546. 1899. margins smooth, reddish brown near the apex, the microglochin (Wahlenb.) Spreng. var. fuegina orifice obliquely truncate. Achenes trigonous with (Kük.) Macloskie, Reports Univ. Exped. narrowly oblong sides, 2.5-3.4 mm long, 0.5-0.9 mm Patagonia, 1896–1899, 8(5): 271. 1904. Carex wide. Rachilla 5-9 mm long, narrowly long-subulate, microglochin Wahlenb. var. oligantha (Boott) somewhat curved, exserted up to 3.5 mm beyond Kük., in A. Engler (ed.), Pflanzenr., 4(20), 38: orifice of perigynium. Stigmas 3; style usually 109. 1909. Type: “Fuegia, Orange Harbor,” slightly exserted, continuous with achene. Anthers [CHILE: Región XII (Magallanes y de la 1.6-2.4 mm long. Antártica Chilena), Prov. Antártica Chilena: In the Fuego-Patagonian region, Carex Isla Hoste, Península Hardy, Bahía Orange, 55° camptoglochin grows primarily in persistently wet, 31’S Lat., 68° 03’W Long.,] 1838–1842, Wilkes base-poor “bogs” (most of them probably more Explor. Exped. s.n. (holotype: K; isotypes: GH, appropriately called poor fens), where Sphagnum US). Figs. 1, 3. hummocks, often dominated by S. magellanicum Brid., are well developed. In these Sphagnum- Plant perennial, vegetative and flowering shoots dominated bogs, the wiry, branched stems of C. arising from dormant buds in axils of dried leaves of camptoglochin often grow nearly vertically, exten- prolonged, branched, decumbent old stems, 0.9-1.4 ding upward to the top of moss-laden hummocks, mm thick; roots produced at lower nodes of old, with the roots seldom coming in contact with mine- prostrate stems; fertile culms 3-34 cm tall, very ral soil. Other typical species in these bogs include, obscurely trigonous, smooth, with glabrous, among many others, Carex magellanica Lam., reddish brown basal sheaths. Leaves 2-8; blades 1- Carpha alpina R. Br., Cortaderia pilosa, Donatia 13 cm long, 0.4-0.8 mm wide, slightly channelled fascicularis J. R. Forst. & G. Forst., Juncus except near the apex where flattened to more or less scheuchzerioides Gaudich., Marsippospermum terete, glabrous, the margins smooth (except grandiflorum (L.f.) Hook. f., Rostkovia slightly scaberulent near the apex), tips blunt; inner magellanica (Lam.) Hook. f., Tetroncium band of sheaths glabrous, pale reddish brown or magellanicum Willd. and the bryophytes reddish-brown-dotted; ligules 0.2-0.4 mm long, 0.5- Calliergon sarmentosum (Wahlenb.) Kindb. and 0.8 mm wide, white-hyaline, the free portion reddish 201 Darwiniana 41(1-4). 2003

Drepanocladus fluitans (Hedw.) Warnst. Carex Hambre, prope Rancho Hambre, 54° 45’S, 67° 54’W, 140 camptoglochin is also known to grow in mossy m s.m., 16-I-1970, Roivainen 1950 (H: 2 sheets, RNG); mats along rivers and in wet meadows, the latter Estancia Harberton, 54° 50’S, 67° 16.5’W, ca. 300 m often at or above tree line. On the Falkland Islands s.m., 22-I-1968, Moore 1602 (C, RNG); Bahía Moat, Estancia Moat, Moat Lake, 54° 57’S, 66° 46’W, 27-I- (Islas Malvinas), where this species is rare, it occurs 1968, Moore 1705 (H, RNG); Estancia Moat, 10-II- from near sea level to about 150 m s.m., growing in 1967, Goodall 678 (NA, RNG); Lapataia, Laguna Negra, “peat bogs” and often in association with W of Río Lapataia, 2-III-1970, Goodall 2417 (BAB, NA, Rostkovia magellanica and Juncus scheuchzerioi- RNG); Río Rancho Lata, near Mt. Moore, Bridges trail, des (Moore, 1968). 1600 ft., 7-II-1971, Goodall 3340 (NA, RNG); Río In northern South America C. camptoglochin Valdez, Bridges track, 1200 ft., 9-II-1971, Goodall 3400 occurs in páramo, where it grows from about 3500 to (NA, RNG); Preto’s sawmill on Ruta 0, Río Remolino, 4200 m s.m. In both Colombia and Ecuador, this Remolino track, 500–1800 ft., 12-II-1971, Goodall 3418 species is most often recorded from persistently wet (NA, RNG); NE of Río Tristen, 28-IV-1974, Goodall 4812 (RNG); Tierra Mayor Valley, 23-III-1974, Goodall places, such as boggy depressions, swamps, slo- 4796 (BAB, NA, RNG); Monte de las Carpas, valley NE ping bogs, swales, and lakeshores, most of which of Monte Olivia, 6-III-1973, Goodall & Prosser 4673b are dominated by hydric mosses and cushion (RNG); N of Almirante Brown, 50 m s.m., 25-I-1979, plants; however, it has also been recorded from Lægaard 13289 (AAU, NY); Isla de los Estados, Bahía boggy depressions dominated by Espeletia. Unlike Franklin, desde Caleta le Croix hacia Crossley, 16-XII- the similar-appearing C. microglochin, which is well 1999 (fl.), Biganzoli 744 (SI); Isla de los Estados, Bahía known from grassy páramo, C. camptoglochin Franklin, desde Caleta le Croix hacia Crossley, 16-XII- seldom (if ever) grows in areas dominated by bunch 1999 (fr.), Biganzoli 746 (SI); Isla de los Estados, Puerto grasses, such as Calamagrostis. Abrigado, 54° 46’S, 64° 09’W, 16-II-1934, Castellanos s.n. (BA 12810, 2 sheets); Isla de los Estados, XII-1932, Notably, several high Andean wetlands in Castellanos 264 (SI). southern Peru, Bolivia, and northwestern Argentina CHILE. Región X (de Los Lagos) Prov. Osorno: and adjacent Chile have recently been intensively Cordillera de la Carpa, ca. 950 m s.m., 1-3-II-1958, studied (Ruthsatz, pers. comm.), but C. Eyerdam 10585 B (F, NY, UC, US). Región XI (Aisén camptoglochin was not recorded from any of these del Gral. Carlos Ibáñez del Campo) Prov. Aisén: Parque sites. While under collecting is a possibility, more Nac. Laguna San Rafael, Istmo de Ofqui, 46° 42’S, 74° likely the absence or paucity of extensive tracts 06’W, 3-4-II-1988, Pisano 6317 (HIP, MIN); Isla dominated by hummock-forming mosses, as is Rivero, Puerto Concell, 45° 36’S, 74° 11’W, 29-XI-1968, characteristic of bofedales (this wetland type is Barrett s.n., Hero No. 38–21 (NA, RNG). Región XII discussed in more detail under C. microglochin (Magallanes y de la Antártica Chilena) Prov. Antártica below), may play no small role. Lastly, it is worth Chilena: Isla Navarino, Puerto Toro, 17-I-1971, Pisano noting that C. camptoglochin is unknown, thus far, 3452 (RNG); Isla Navarino, Puerto Toro, 2-III-1976, Dollenz 383 (GH, HIP, NA); Isla Navarino, Bahía from north of Colombia. Windhond, 5-6-III-1976, Dollenz 417 (HIP); Islas Wollaston: Isla Bayly, Surgidero Romanche, 55° 37’S, Representative specimens of 67° 33’W, 19-II-1980, Pisano 5056 (HIP, RNG); Isla Carex camptoglochin Hoste, Caleta Awaiakirrh, 23-I-1982, Pisano 5469 (GH, ARGENTINA. Tierra del Fuego, Antártida e Islas HIP, RNG); Smoke Island, 54° 43’S, 71° 07’W, 18-V- Atlántico Sur. Dpto. Islas del Atlántico Sur: Islas 1882, Spegazzini s.n. [227 of Kükenthal, 1909: 109] Malvinas (Falkland Islands), East Falkland Island (Isla (CORD 14029, LP s.n.). Prov. Magallanes: Península de Soledad): Port Stanley, Mount William, 28-II-1904, Brunswick, Cerro Mirador, camino al Club Andino, 8-II- Birger s.n. (S); Stanley, 1-I-1902, Skottsberg s.n. (S: 2 1979, Dollenz 608 (HIP, NA); Fiordo Silva Palma, An- sheets); Stanley, valley N of Two Sisters, 250-300 ft., 14- gostura Titus, 8-I-1973, Pisano 3780 (HIP, MO, NA, I-1968, Engel s.n. (H-1459345); West Falkland Island RNG); Laguna El Parrillar, 53° 25’S, 71° 17’W, 7-II- (Isla Gran Malvina): Hill Cove, base of French Peaks, ca. 1983, Pisano 5730 (GH, HIP); Isla Santa Inés, Fiordo 500 ft., 14-II-1964, Moore 873 (LP, RNG). Dpto. Nevado, Bahía Havergal, 53° 31’S, 72° 46.6’W, 26-II- Ushuaia: Ushuaia, 275 m s.m., 25-II-1940, Santesson 1990, Pisano & Henríquez 6698 (HIP, MIN); Isla 554 (S); Ushuaia, 245 m s.m., 11-III-1902, Skottsberg s.n. Clarence, Estero Staples, 15-II-1929, Roivainen s.n. (H: 2 (S); Montes Martiales, 350 m s.m., 21-XII-1969, sheets); Isla Vidal Gormaz, Seno Nantuel, Bahía María Roivainen 1252 (H: 2 sheets, RNG); Montes Martiales, Angélica, 51° 53’S, 74° 41’W, 3-II-1976. Dollenz et al. 9-I-1960, Correa & R. L. Pérez Moreau 1848 (BAB); Río s.n., T.B.P.A. 1306 (BAB). Prov. Última Esperanza:

202 G. A. WHEELER & E. R. GUAGLIANONE. Notes on South American Carex: C. camptoglochin and C. microglochin

Parque Nac. Torres del Paine, Cerro Quixote, W of Lago Carex microglochin Wahlenb., Kungl. Svenska Grey, 51° 00’S, 73° 25’W, III-1976, Steele 84 (RNG); Vet.–Akad. Handl. 24: 140. 1803. Figs. 2, 4. Fiordo Peel, Río Murtillar, 50° 27’S, 73° 37’W, 20-XI- 1985, Pisano 5952 (HIP, MIN, RNG); Fiordo Falcon, Detailed descriptions of Carex microglochin barranca en costa E, 49° 37’S, 73° 50’W, 21-XII-1988, already exist in the literature, such as that provided Pisano 6509 (HIP, MIN); Isla Piazzi, 51° 02’S, 74° by Mackenzie (1935: 424), therefore one is not given 00’W, 16-I-1976, Eskuche s.n., T.B.P.A.-FIT. 704 here. Also see Mackenzie for synonymy and type (BAB); Puerto Bella Vista, 51° 30’S, 73° 15’W, 9-XII- information, and Luteyn (1999: 105). 1979, Roig et al. s.n., T.B.P.A.-FIT. 5280 (BAB, CONC); Isla Piazzi, Caleta Ocasión, Abra Leackey’s The South American populations of C. Retreat, 51° 44’S, 74° 01' W, 80 m s.m., 19-I-1976, microglochin grow primarily in moist to wet sites Dollenz et al. s.n., T.P.B.A. 1080 (BAB, RNG); Isla that are either deficient in mosses or at least in Rennel Norte, Canal Smyth, 51° 54’S, 74° 12’W, 250 m places where Sphagnum hummocks are poorly- s.m., 28-I-1976, T.B.P.A. 1221 (HIP, RNG)); Isla Virtu- des, Canal Elías, Puerto Virtudes, 51° 33’S, 74° 54’W, developed. For instance, high Andean bofedales in 12-II-1976, Dollenz et al. s.n., T.B.P.A. 1511 (BAB, Chile, Argentina, and Bolivia do not harbor many HIP, RNG); Seno Última Esperanza, Puerto Toro, Río mosses; rather, these bofedales are predominantly Serrano, 51° 25’S, 73° 05’W, 16-I-1977, Moore & Pisano formed by cushion-forming Juncaceae (Ruthsatz, s.n., T.B.P.A. 1732 (BAB, HIP, RNG); Seno Última per. comm.), with diminutive carices, such as C. Esperanza, Puerto Toro, Río Serrano, N of Vent. microglochin and C. ruthsatzae G. A. Wheeler, Balmaceda, 51° 25’S, 73° 05’W, 18-I-1977, Moore & growing in wet soil between cushion-forming Pisano s.n., T.B.P.A. 1788 (BAB, HIP); Seno Última vascular plants, such as Oxychloë andina Phil. and Esperanza, Lote Sanchez, costa E Río Serrano, 51° 26’S, Distichia muscoides Nees & Meyen (Wheeler, 73° 05’W, 26-I-1977, T.B.P.A. 2037 (BAB, HIP, RNG). 2002). Similarly, near Cochrane, in Capitán Prat COLOMBIA. Caldas. Cordillera Central, Páramo del Province (Región XI, Aisén), Chile, C. Quindio, 3700–4200 m s.m., 15-20-VIII-1922, Pennell & Hazen 9961 (GH, NY). Cauca. Volcán Puracé, Laguna de microglochin grows in mires around lakes, such as San Rafael, aprox. 2 km al S de la Laguna, 3450 m s.m., 31- Laguna Elefantita and nearby Laguna Tamange, III-1972, Cleef 2661 (NY). where Sphagnum bogs are absent. These mires are ECUADOR. Carchi. Road Tulcán–Maldonado, 34– dominated by Poaceae and Cyperaceae, primarily 36 km from Tulcán, at base of Volcán , 00° 47’N, Cortaderia pilosa and Schoenus andinus (Phil.) H. 77° 57’W, 3950 m s.m., 2-VIII-1976, Øllgaard & Balslev Pfeiffer, with diminutive carices, such as C. 8346 (AAU, CTES, NY); Road Tulcán–Maldonado, microglochin and C. capitata L., growing in the approx. 43 km from Tulcán, SW slopes of Volcán Chiles, lower, herbaceous layer; several mosses also occur 00° 48’N, 77° 59’W, 3740 m s.m., 4-VIII-1976, Øllgaard in the ground layer, but not Sphagnum (Vogel, per. & Balslev 8539 (AAU); El Angel–San Gabriel km 8 along comm.). In San Juan Province, Argentina, C. old road, páramo of Hac. El Consuelo, 00° 37’N, 77° microglochin is known to grow among vegetation 52’W, 3475–3625 m s.m., 15-II-1992, Lægaard 101319 bordering fast-flowing streams, from about 3400- (AAU); Páramo del Angel, km 18 along road El Angel– Tulcán, páramo del Voladero, around large lakes near the 3900 m s.m. These moist to wet riparian margins, pass, 00° 41’N, 77° 53’W, 3650–3750, 9-VIII-1990, which often harbor scattered plants of Acaena Øllgaard 98087 (AAU). Chimborazo. Road ca. 10 km magellanica (Lam.) Vahl. and A. pinnatifida Ruiz & NE of Alao, at Cuspipaccha, 01° 48’S, 78° 26’W, 3550– Pavón, frequently have in the ground layer the 3600 m s.m., 6-V-1982, Øllgaard et al. 38121 (AAU, cushion plants Oxychloë simulans Barros, Azorella QCA); About 15 km NE of Aloa on trail to Huamboya, at trifurcata (Gaertn.) Pers., and A. crypthanta (Clos) the bridge over Río Alao, 01° 52’S, 78° 28’W, 3500 m Reiche, these often interspersed with other s.m., 4-I-1988, Molau et al. 2329 (AAU). Napo. herbaceous plants growing in mineral soil, such as Llanganati, N-facing slope towards Río Golpe, just N of C. microglochin. Mosses are also usually present, Chosa Aucacocha, 01° 08’S, 78° 18’W, 3600 m s.m., 16- V-1982, Øllgaard et al. 38665 (AAU). Pichincha. but Sphagnum is absent or at least not dominant in Mountain ridge on right side of Río Bolteado, approach the ground layer (Kiesling, per. comm.). Finally, in from the west to Cerro Saraurcu, 00° 03’S, 77° 58’W, northern South America, such as in Ecuador, C. 3800 m s.m., 31-VIII-1995, Sklenar & Kosteckova 1553 microglochin grows in páramo above 3500 m s.m., (AAU). Tungurahua. Cordillera de Llanganates, near primarily in moist to wet sites dominated by grasses, Lake Yanacocha, 3600 m s.m., 16-XI-1939, Asplund 9736 such as Calamagrostis, as well as along the wet (NY). 203 Darwiniana 41(1-4). 2003 margins of streams. In these habitats Carex 1968, Goodall 1127 (NA); Valle Lasifashaj, Las Cotorras, microglochin grows in the herbaceous ground about 20 km ENE of Ushuaia, 200 m s.m., 6-II-1940, layer in mineral soil. Santesson 421 (S). Tucumán. Dpto. Tafí: Peñas Azules, In southern Patagonia and Tierra del Fuego, C. Casa de Piedra, San José, 3150 m s.m., XII-1931, Schreiter 6989 (LIL). microglochin is most frequently recorded on herbarium sheet labels as growing in mountain BOLIVIA. Cochabamba. Prov. Quillacolla: Bella “meadows” (often called vegas or mallines), depre- Vista, Cotopata, Laguna Marquina, 4270 m s.m., 3-II- 1999, Ruthsatz & Budde 10259 (MIN). La Paz. Prov. ssions in grasslands, along river banks and, less Murillo: La Paz, Río Palcoma (Río Hampaturi), 4370 m frequently, in swamps. These moist to wet sites, s.m., 15-II-1999, Ruthsatz & Budde 10345 (MIN); Valle most of which are dominated by grasses and de Chuquiaguillo, Paeocola Pampa, 4200 m s.m., 4-I- sedges, generally harbor several mosses, 1921, Asplund 1945 (UPS). sometimes including Sphagnum species, but the CHILE. Región IV (Coquimbo) Prov. Limarí: mosses seldom dominate or form well-developed Bocatoma Río Hurtado, Seitental, 2650 m s.m., 1-XII- hummocks in these habitats. In this region, like 1990, Ruthsatz 6508 (MIN). Región XI (Aisén del Gral. farther northward in the Andes, C. microglochin Carlos Ibáñez del Campo) Prov. Capitán Prat: NW of seemingly grows in places where its roots and Lago Cochrane, Lago Elefantita, 47° 12’S, 72° 31’W, rhizomes come in contact with mineral soil. It is also 1000 m s.m., 26-II-1999, Vogel 43 (MIN). Región XII noteworthy that in the Northern Hemisphere C. (Magallanes y de la Antártica Chilena) Prov. Tierra del Fuego: Lago Blanco (E side), Cerro Cuchillo (E side), 54° microglochin is often reported as growing in moist, 05’S, 68° 59’W, 12-III-1968, Moore 2222 (H); Porvenir, calcareous soil (e.g., Mackenzie, 1931–1935; 2-I-1941, Santesson 1550 (S). Prov. Última Esperanza: Damman, 1964), as well as in sites “where the total Lago Balmaceda, 4-II-1938, Kalela 2031 (H). plant cover is usually less than 50 percent” (Jermy ECUADOR. Cotopaxi. Zumbahua-Pujilí km 39, 00° et al., 1982: 210). In the Northern Hemisphere, to my 53’S, 78° 47’W, 3750-3800 m s.m., 4- IV-1992, Lægaard knowledge (G.A.W.), this species has not been 102093 (AAU, QCA); Latacunga–Quevedo road, 28.5 reported from well-developed Sphagnum bogs. km W of Latacunga, 00° 52’S, 78° 43’W, 3780 m s.m., 4- IV-1992, Luteyn & Quezada 14392 (AAU, QCA); Representative specimens of Carex microglochin Latacunga-Quevedo road, 18 km W of Pujilí, 3700 m s.m., ARGENTINA. Jujuy. Dpto. Susques: subida de Cho- 28-XI-1980, Balslev 1057 (NY). rrillos, 4100 m s.m., II-1980, Cabrera et al. 31756 (SI). PERU. Huánuco. Prov. Lauricocha: San Miguel de La Rioja. Famatina, camino a Alto Blanco, 20-I-1928, Cauri, Laguna Taulicocha (E side), 10º 20’S, 76º 44’W, Castellanos s.n., BA 28/33 a (SI). Salta. Dpto. Cachi: 3910 m s. m., 9-VIII-2002, Salvador 434 (MIN); ídem, Cerro Negro, 3710 m s.m., 5-I-1994, Charpin & Lazare Salvador 438 (MIN); Ídem, Añaspampa bog; 10º 21’S, 24055 (G, MIN); Dpto. Santa Victoria: Paso de Lizoite, 76º 45’W, 3915 m s.m., 10-VIII-2002, Salvador et al. 22° 16’S, 65° 11’W, 4355 m s.m., 7-III-1997, Ruthsatz & 510. (MIN). Lima. Prov. Cañete: Cañete valley, above Erschb 9822 (MIN). San Juan. Dpto. Iglesia: Quebrada Hortigal near Madean, 12° 57’S, 75° 46’W, 3600 m s.m., del Agua Negra, El Arenal, 30° 20’S, 69° 40’W, 3700– 7-III-1987, Brandbyge 223 (AAU). 3900 m s.m., 24-II-1992, Kiesling et al. 7996 (SI); Que- Some selected sites from the Northern Hemisphere. brada Agua Negra, Arroyo Pircas, 3900 m s.m., 20-I- 1995, Ruthsatz 8985 (MIN); Cordillera de Colangüil, CANADA: Alberta. Banff National Park, near Banff, Quebrada de la Vicuñita, 3400 m s.m., 26-I-1995, Kiesling 1-VII-1915, Hunnewell 3965 (MIN); Manitoba. vicinity et al. 8707 (SI). Santa Cruz. Dpto. Lago Argentino: Lago of Churchill, 58° 46’S, 74° 10’W, 18-VII-1956, Schofield San Martín, Bahía del Depósito, 8-III-1905, Dusén 6040 & Crum 6821 (MIN); Newfoundland. Straits of Belle (BAF, S); Brazo Norte: valle de La Cristina, pie occiden- Isle, Capstan Point, Flower Cove, 28-VII-1924, Fernald tal cerro Cuchillo, 970 m s. m., abundante en el borde del et al. 26403 (MIN). mallín, 4-II-1953, Vervoorst 4406 (LIL). Tierra del Fue- NORWAY: Dovre, Kongsvold, 18-VII-1885, Olsson go, Antártida e Islas del Atlántico Sur. Dpto. Río s.n. (SI-2923); Ringvatsö Island, North of Tromsö, 10- Grande: Estancia La Criolla, 19-XII-1972, Goodall 4414 VI/10-VII-1896, Notø 31 (SI). (A, BAB, MO, NA, RNG); Río de la Turba, Estancia La SWITZERLAND: VII-1875, Colemann s.n. (SI). Marina, 54° 18’S, 68° 30’W, 23-II-1968, Moore 2027 (C, UNITED STATES OF AMERICA: Alaska. Arctic RNG). Dpto. Ushuaia: Estancia Harberton, Campo Lagu- North Slope, Anaktuvuk Pass, 68° 20’N, 151° 25’W, nas Abajo, Harberton Swamp, 54° 51’S, 67° 20’W, 5-I- 2000 ft, 7-VIII-1951, Spetzman 3976 (MIN); Colorado. 1968, Moore 1343 (BAB, H, MO, RNG); Estancia Denver, Three Cabins, 12000 ft., 8-XI-1919, Clokey Harberton, Harberton Swamp and Fox Trap Hill, 5-I- 3342 (BAF, MIN).

204 G. A. WHEELER & E. R. GUAGLIANONE. Notes on South American Carex: C. camptoglochin and C. microglochin

Key to the three South American members of Carex sect. Leucoglochin 1. Apex of rachilla tipped with a hook-like appendage; perigynia ascending to somewhat wide spreading, but not deflexed, at maturity ...... C. kingii 1. Apex of rachilla lacking a hook-like appendage; perigynia wide spreading or, more often, strongly deflexed at maturity ...... 2 2. Pistillate scales as long as or longer than perigynia, apex acute to acuminate or smooth-awned, reddish brown; staminate scales 2-3 per spike; perigynium long-stipitate, when deflexed the stipe appearing inverted U-shaped, beak reddish brown near apex; perigynia 2–5 (most often 3 or 4) per spike; plants with stems becoming decumbent and sending up new stems from dormant buds in old leaf axils ...... C. camptoglochin 2. Pistillate scales shorter than perigynia, apex obtuse to subacute, brownish; staminate scales 5-6 per spike; perigynium short-stipitate, when deflexed appearing subsessile on the rachis, beak stramineous or brownish at apex; perigynia 3-12 per spike; plants rhizomatous ...... C. microglochin

ACKNOWLEDGMENTS Damman, A. W. H. 1964. Key to the Carex species of We want to thank R. Kiesling (Instituto de Botánica Newfoundland by vegetative characteristics. Can. Darwinion, Argentina), J. Luteyn (New York Botanical Dept. Forestry Publ. 1017. Garden, U.S.A.), B. Ruthsatz (University of Trier, Guaglianone, E. R. 1996. Carex. In: F. O. Zuloaga & O. Germany), F. Salvador (Universidad de Alicante, Spain) Morrone (eds.), Catálogo de las Plantas Vasculares and A. Vogel (University of Munster, Germany) for de la República Argentina. I. Monogr. Syst. Bot. distribution and habitat information on Carex Missouri Bot. Gard. 60: 133-146. microglochin and also some of them for providing Hermann, F. J. 1970. Carices of the Rocky Mountains duplicate specimens; D. Simpson (Royal Botanic and Colorado Basin. U.S.D.A. Handb. No. 374. Gardens, Kew) for sending an image of the type of Carex Holmgren P. K., Holmgren N. H. & Barnett, L. C. 1990. oligantha Boott; A. Vizinis for the fine drawings; A. Index Herbariorum. Part 1: The Herbaria of the Garbini (Instituto de Botánica Darwinion) for technical World. New York Botanical Garden. New York. assistance; A. Cholewa (University of Minnesota) for Jermy, A. C., Chater, A. O. & David, R. W. 1982. Sedges arranging loans and providing other herbarium services of the British Isles. B.S.B.I. Handb. No. 1, London. and assistance; and also to the directors and curators of Jørgensen, P. M. & Ulloa Ulloa, C. 1994. Seed plants of the following herbaria for the in-house study or loan of the High Andes of Ecuador: a checklist. AAU specimens: A, AAU, BA, BAB, BAF, C, CONC, Reports 34: 1-443. CORD, CTES, F, G, GH, H, HIP, K, LIL, LP, MIN, Kükenthal, G. 1909. Cyperaceae: Caricoideae. In: A. MO, NA, NY, QCA, RNG, S, SI, UC, and UPS (cited Engler (ed.), Pflanzenr. 4. 20, Heft 38: 1-824. according to Holmgren et al., 1990). Wilhelm Engelmann, Leipzig. Kukkonen, I. 1970. Vegetative anatomy of Carex LITERATURE CITED microglochin Wahlenb. and Carex camptoglochin Krecz., Bot. J. Linn. Soc. (London) 63, Suppl. 1: Barros, M. 1947. Cyperaceae: Scirpoideae, Rhynchospo- 137-145. roideae, Caricoideae. In: H. R. Descole (ed.), Gen. Kreczetowicz, V. I. 1937. Cyperacearum novitates. Bot. Sp. Pl. Argent. 4(2): 259-539. Mater. Gerb. Bot. Inst. Komarova Akad. Nauk S. S. ——. 1969. Cyperaceae. In: M. N. Correa (ed.), Flora S. R. 7: 27–37 Patagónica, Typhaceae a Orchidaceae (excepto Luteyn, J. L. 1999. Páramos: A checklist of plant Gramineae). Colecc. Ci. Inst. Nac. Tecnol. Agropecu. diversity, geographical distribution, and botanical 8(2): 38-92. literature. Mem. New York Bot. Gard. 84: 1-278. Boott, F. 1867. Ill. Carex, 4: 127-233. L. Reeve and Co., Mackenzie, K. K. 1931–1935. Cyperaceae: Cariceae. London. North Amer. Flora 18(1-7): 1-478. New York Chater, A. O. 1994. Carex. In: Davidse, G., M. Souza & Botanical Garden, New York. Parts 1-3, 1931; 4-7, A. O. Chater (eds.). Fl. Mesoamericana 6: 464-473. 1935. Universidad Nacional Autónoma de México, Méxi- Marticorena, C. & Quezada, M. 1985. Catálogo de la flora co, D. F.; Missouri Bot. Gard., St. Louis and the vascular de Chile. Gayana, Bot. 42: 1-157. Natural History Museum, London. Moore, D. M. 1968. The vascular flora of the Falkland Islands. Sci. Rep. Brit. Antarct. Surv. 60: 1-202.

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——. 1974. Catálogo de las plantas vasculares nativas de Reznicek, A. A. & Catling, P. M. 1986. Vegetative shoots Tierra del Fuego. Anales Inst. Patagonia 5: 105–121. in the taxonomy of sedges (Carex, Cyperaceae). ——. 1983. Flora of Tierra del Fuego. Anthony Nelson Taxon 35: 495-501. Ltd. and Missouri Botanical Garden. ——. 1990. Evolution in sedges (Carex, Cyperaceae). —— & Chater, A. O. 1971. Studies of bipolar disjunct Canad. J. Bot. 68: 1409-1432. species. I. Bot. Notiser 124: 317-334. Roivainen, H. 1954. Studien über die Moore Feuerlands. Mora-Osejo, L. E. & Rangel, O. 1983. Una nueva Ann. Bot. Soc. Zool. Bot. Fenn. Vanamo 28: 1-205. Cyperaceae de la Sierra Nevada de Santa Marta (Co- Wheeler, G. A. 2002. A new species of Carex section lombia) y consideraciones fitogeográficas y Abditispicae (Cyperaceae) from South America and sinecólogicas sobre Carex L. Academia Colombiana additional notes on the section. Darwiniana 40: 191- 15: 13-21. 198. Raven, P. H. 1963. Amphitropical relationships in the floras of North and South America. Quart. Rev. Biol. Original recibido el 24 de septiembre de 2002; acepta- 38: 151-177. do el 12 de junio de 2003.

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